Exxon Valdez Oil Spill Restoration Project Final Report
Recovery Monitoring of Pigeon Guillemot Populations in Prince William Sound, Alaska
Restoration Project 94173 Final Report
D. Lindsey Hayes
U.S.Fish and Wildlife Service 1011 East Tudor Road Anchorage,Alaska 99503
May 1995 Recovery Monitoring of Pigeon Guillemot Populations in Prince William Sound. Alaska
Restoration Project 94173 Final Report
Studv Historv: The field work for Restoration Project 94173 was conducted during the summerof 1994. Previous related projects have been funded by the Trustee Council. Bird Study Number 9 (Oakley and Kuletz 1994),begun in 1989 immediately after theoil spill, compared various population and reproductive parameters of pigeon guillemots before (Oakley and Kuletz 1979, Kuletz 1981, 1983, Oakley 1981) and after the spill. Also, Project 93034, an extensive survey of pigeon guillemot colonies throughout Prince William Sound, was conducted during the summer of 1993 (Sanger andCody 1994).
Abstract: The population of pigeon guillemots in Prince William Sound decreased from about 15,000 (1970's) to about 5,000 (present). Some local populations were affected by the TIV Exxon Valdez oil spill in 1989, but there is evidence suggesting the Sound-wide population was already declining. In 1994, we monitored 22 guillemot nests on Naked Island (NI) and24 on Jackpot Island (JI) from egg stage through fledging. Hatching success was 0.90 (n = 39, NI) and 0.80 (n = 46, JI); fledging success was 0.51 (n = 35, NI) and 0.76 (n = 37, JI). On NI, predation was the cause of numerous nesting failures. The most likely predators were mustelids. Predation on eggs by corvids was also suspected. Although predation was infrequent or nonexistent on JI, abandonment of eggs was high. Sand lance accounted for about 1% and 8% of prey items delivered to guillemot chicks at JI andNI in 1994; before the spill, sand lance were the singlemost important component of chick diet at NI. Gadids were more prevalent in the diet of guillemot chicks at NI in 1994 (ca. 30%) than in 1979-1981 (< 7%). Changes in the relative proportions of benthic and schooling fish in the dietof guillemot chicks might represent a key change in the ecosystem that is affecting other species of marine birds and mammals in the Sound.
Kev Words: Alaska, Cepphus columba, foraging ecology, Jackpot Island, Naked Island, oil spill, pigeon guillemot, population, Prince William Sound, predation, productivity, TIV Exxon Valdez
Citation: Hayes, D.L. 1995. Recovery monitoring of pigeon guillemot populations in Prince William Sound, Alaska, Exxon Valdez Oil Spill Restoration Project Final Report (Restoration Project 94173), U.S. Fish and Wildlife Service, Anchorage, Alaska. TABLE OF CONTENTS
LISTOFTABLES ...... iii
LIST OFFIGURES ...... iv
ACKNOWLEDGMENTS ...... vi
EXECUTIVESUMMARY ...... vii
INTRODUCTION ...... 1
OBJECTIVES ...... 2
METHODS ...... 2 StudyArea ...... 2 Selection of StudySites ...... 3 Censusing:Population and Colony Attendance ...... 3 NestSites and Monitoring ...... 4 Bandingand Morphometrics ...... 4 Nesting Chronology ...... 5 Productivity ...... 5 Predation ...... 6 Chick Growthand Fledging Weights ...... 6 Chick Provisioning and Diet ...... 7 Identificationand Sampling of Fish Types ...... 8 DataAnalysis ...... 8
RESULTS ...... 8 Censusing: Population and Colony Attendance ...... 8 Nesting Chronology ...... 9 Productivity ...... 10 Predation ...... 10 Chick Growth and Fledging Weights ...... 11 Chick Provisioning and Diet ...... 11 Fish Types Caught in Traps ...... 12
DISCUSSION ...... 12 Censusing:..Population and Colony Attendance ...... 12 Productivity ...... 13 Predation ...... 14 OtherCauses of NestingFailure ...... 16 ChickGrowth andFledging Weights ...... 16 ChickProvisioning ...... 18
i Foraging ...... 19 Chick Diet ...... 19
CONCLUSIONS ...... 20
LITERATURE CITED ...... 22
APPENDIX ...... 66
.. 11 LIST OF TABLES
Table 1. Counts of pigeon guillemots during June censusesat Naked, Peak,Storey, Smith, and Little Smith Islands, Prince William Sound, Alaska, before and after the T/V Exxon Vuldez oil spill ...... 29
Table 2. Number of active pigeon guillemot nests at five colonies on the western sideof Naked Island, Prince William Sound, Alaska, before and after theT/V Exxon Valdez oilspill ...... 30
Table 3. Growth rates of pigeon guillemot chicks raised at Naked Island, Prince William Sound, Alaska, before and after the T/V Exxon Vuldez oil spill ...... 31
Table 4. Fledging weights of pigeon guillemot chicks raised at Naked Island, Prince William Sound, Alaska, before and after the T/VExxon Vuldez oil spill ...... 32
... 111 LIST OF FIGURES
Figure 1. Locations investigated during the 1994 field season as potential study sites forpigeon guillemots breeding in Prince William Sound, Alaska ...... 35
Figure 2. The Naked Island complex, where pigeon guillemots were censused in 1978-1980 and in 1989-1994 .... 37
Figure 3. Location of pigeon guillemot colonies on Naked Island monitored during the 1994 field season ...... 39
Figure 4. Attendance patterns of pigeon guillemots at two colonies on Naked Island during the incubation period ...... 41
Figure 5. Attendance patterns of pigeon guillemots at two colonies on Naked Island during the early hatching period ...... 43
Figure 6. Nesting chronology of pigeon guillemots on Naked Island during the 1994 breeding season ...... 45
Figure 7. Nesting chronology of pigeon guillemots on Jackpot Island during the 1994 breeding season ...... 47
Figure 8. Comparison of body mass vs. wing length measurements of pigeon guillemot chicks from Naked and Jackpot Islands ...... 49
Figure 9. Temporal distribution of food deliveries by adult pigeon guillemots at several colonies on Naked Island during the 1994 breeding season ...... 51
Figure 10. Temporal distribution of food deliveries by adult pigeon guillemots at Jackpot Island during the 1994 breeding season ...... 53
Figure 11. Temporal distribution of food deliveries to each of four guillemot nests at the Tuft colony on Naked Island ...... 55
Figure 12. Temporal distribution of food deliveries to each of three guillemot nests at the Nomad colony on Naked Island ...... 57
Figure 13. Temporal distribution of food deliveries to each of four guillemot nests on Jackpot Island ...... 59
iv Figure 14. Bathymetry of the Naked Island area ...... 61
Figure 15. Bathymetry of the Jackpot Island area ...... 63
Figure 16 . Comparison of prey types delivered by adult pigeon guillemots at Naked and Jackpot Islands ...... 65
V ACKNOWLEDGMENTS
I thank Mary Cody, Kirk Lenington, John Maniscalco, Bev Short, and Ed Vorisek for help in the field, and Mark Esslinger and Burt Prattefor help with logistics. The crew of the Alaskan Gypsy and the marbled murrelet field team helped with the capture of adult pigeon guillemots. The U.S. Forest Service granted us permission to use Naked Island as our base of operations while in the field. I also thank Debbie Flint for making the maps and Mary Blakefor typing the tables. Bryan Manly suggested the use of certain statistical analyses for some of the data andperformed the randomization analysis of the growth data. Discussions with George Divoky, David Irons, Kathy Kuletz, and Dan Roby regarding various aspects of this project have been most helpful. The original manuscript of this report hasbeen greatly improved through many helpful suggestions by Tony DeGange.
vi EXECUTIVE SUMMARY
The pigeon guillemot (Cepphus columba) is a cavity-nesting, pursuit-diving seabird that forages in nearshore waters. Guillemots are common, easily observed birds in parts of coastal Alaska. Between the early 1970's and the early199O's, the population of pigeon guillemots in Prince William Sound (PWS) decreased from about 15,000 to less than 5,000. Over 600 carcasses of pigeon guillemots were recovered in the four months following the T/V Exxon Vuldez oil spill in March of 1989, and these probably represent only a fraction (ca. 10-30%)of the actual number killed, Although some local populations of guillemots were affected by the spill, there is evidence suggesting that theSound-wide population was already declining. We studied the breeding and feedingecology of pigeon guillemots nesting on two islands in the western part of PWS during the 1994 field season. Our objectives were to examine the effects of food availability and diet, predation, and adult survival and recruitmenton productivity and numbers of guillemots in different parts of PWS. We marked 51 pigeon guillemot nests on Naked Island (NI) and 37 nests on Jackpot Island (JI). Of these nests, we monitored 22 on Naked I. and 24 on Jackpot I. from the egg stage through fledging. Hatching success was 0.90 (n = 39, NI) and 0.80 (n = 46, JI); fledging success was 0.51 (n = 35, NI) and 0.76 (n = 37, JI). Chicks raised on Jackpot I. grew at a faster rate and fledged at significantly greater weights than chicks raised on Naked I. On Naked I., predation during thechick stage was the causeof numerous nesting failures (8 of 35 nests, or 23%). Decapitated carcasses of chicks found near depredated nests suggest that the predators were mustelids. River otters (Lutra canadensis) were often seen around the guillemot colonies, and were the most likely predator, although mink might also have been responsible. Predation on eggs by corvids was also suspected. Although we saw the same species of potential avian and mammalian predators on or around Jackpot I.,we found no evidence of predation. However, the abandonment of eggs on Jackpot I. was much higher than on Naked I. Depredation of guillemot nests on Naked I. was not common during previous studies, but is now a major factor affecting their reproductive success. Studies at other guillemot colonies have related lowered productivity or emigration to the presence of mammalian predators, especially otters and mink. The adults breeding on Naked I. foraged mostly in the shallow waters surrounding the island. Guillemots breeding on Jackpot I. foraged farther from their colony, apparently because of the lack of shallow water around Jackpot I. Based on their flight directions, these birds presumablyforaged near Jackpot Bay and Icy Bay. There was a marked difference in the proportion of schooling fish in the diet of chicks at the two islands. Sand lance accounted for about 1%and 8% of prey items delivered to guillemot chicks at Jackpot I. and Naked I. in 1994. Before the spill, this species was the singlemost important component of the chick
vii diet at Naked I. Gadids were much more prevalent in the diet of guillemot chicks at Naked I. in 1994 (ca. 30%) than in 1979-1981 (< 7%). There are no historical data for Jackpot I. The apparent decline in the abundanceof sand lance and changes in the relativeproportion of benthic and schooling fish in the diet of guillemot chicks might represent a shift in theecosystem that is affecting other species of marine birds and mammals inPWS.
... Vlll INTRODUCTION
The pigeon guillemot (Cepphus columbu) is a pursuit-diving seabird that forages mostly in nearshore waters about 10-30 m deep (Storer 1952, Ewins 1993). Adults feed primarily on benthic fish and invertebrates, but alsoon schooling fish. Chicks are fed mostly fish. Guillemots nest in small scattered colonies or in solitary pairs in natural cavities alongrocky shorelines. Unlike most other members of the family Alcidae, the pigeon guillemot typically lays a clutch of two eggs. The chicks are semiprecocial, usually spending about 35 to 45 days in the nest. During the daylight hours, they are fed by both parents, which return to the nest with one fish at a time in their bills. The population of pigeon guillemots in Prince William Sound (PWS) has decreased from about 15,000 in the 1970's (Dwyer et al. ND) to less than about 5,000 in the 1990's (Agler et al. 1994, Sanger and Cody 1994). There is some evidence suggesting that this population was indecline before the TI V Exxon Vuldez oil spill in March of 1989 (Oakley and Kuletz 1994). Over 600 guillemot carcasses were recovered after the spill, but this might representonly 10-30% of the actual number killed (Piatt et al. 1990). Based on censuses taken around the Naked Island complex (Naked, Peak, Storey, Smith, and Little Smith Islands), prespill counts (ca. 2,000 guillemots) were roughly twice as high as postspill counts (ca. 1,000 guillemots; Oakley and Kuletz 1994). Also, on Naked I., the relative decline in the numbers of guillemots was greater alongoiled shorelines than along unoiled shorelines (Oakley and Kuletz 1994). King and Sanger (1979)considered the pigeon guillemot to be one of the birds that is most vulnerable to oil spills because of its nearshore foraging habits. Several studies have reported sublethaltoxic effects of oil on marine birds (Peakall et al. 1980, Peakall et al. 1982, 1983 as cited in Oakley and Kuletz 1994). Marked declines in populations of the pigeon guillemot or its congener, the black guillemot (C. grylle) have been attributed to oil pollution (Ainley and Lewis 1974, Asbirk 1978, Ewins and Tasker 1985). At Naked I., adult guillemots delivered fewer schooling fish, particularly sand lance (Ammodytes hexupterus), to their chicks after the spill thanbefore the spill (Oakley and Kuletz 1994). Numerous studies have shown that changes in the availability of prey species can result in widespread reproductive failureof seabirds (Vermeer et al. 1979, Anderson et al. 1982, Springer et al. 1986, Safina et al. 1988, Uttley et al. 1989, Furness and Barrett 1991; but see Burger and Piatt 1990). Predation on eggs and chicks, not important previously on Naked I. (Oakley 1981, Kuletz 19831, was a major factor contributing to the lower reproductive success of guillemots after the spill (Oakley and Kuletz 1994). Studies at other guillemot colonies have related lowered productivity or emigration to the presence of mammalian predators (Asbirk 1978, Petersen 1979, Cairns 1985, Ewins 1985, 1989).
1 Thus, because 1) pigeon guillemots constitute an injured resource, 2) their population has been declining for some time, 3) there has been a marked change in their diet, 4) predation at the nest is more prevalent than in the past, and 5) there exist valuable prespill data for this species in PWS, they have been selected for intensive study. We studied the breeding and feeding ecology of pigeon guillemots nesting on two islands in the western partof PWS and found important differences between the two populations relative to the productivity of the colonies, the foraging habits of adults, diet of chicks, and the levels of predation occurring during the chick stage.
OBJECTIVES
1. Determine if availability offood is limiting reproductive success of guillemots by collecting the following kinds of data: a.Measuring breeding parameters, including phenology, chick growth rates, fledging weights, and reproductive success at colonies on Naked and Jackpot Islands. b. Measuringforaging parameters, including diet and provisioning rates of chicks, and location of foraging areas. 2. Determine if predation on eggs or chicks is limiting reproductive success by measuring relative ratesof predation during theegg and chick stage in different habitats and at different colonies. 3. Determine if adult survival and recruitment are affecting the population of guillemots by banding adults and chicks. METHODS
Study Area Our field season extended from 19 May through 22 August 1994. Our two principal study sites were located on Naked I. and Jackpot I. in PWS (Fig. 1). Naked I. (ca. 3,862 ha) has a maximum elevation of 400 m and is part of a group of three main islands. The bays of Naked I., and the passages between it and the two neighboring islands, Peak and Storey,form an expanse of water that is less than 100 m deep. Jackpot I. (ca. 1.6 ha) has a maximum elevation of about 15 m and is located near the mouth of Jackpot Bay and the southern entranceto Dangerous Passage. Fool I. (ca. 12 ha, Fig. 1) was also a study site initially, but was discontinued later in the season. It lies inWells Passage between Perry and Esther Islands, has a maximum elevation of about 20 m, and is surrounded by deep water on all sides. The shoreline of each of these islands is characterized by low cliffs and cobble or boulder beaches; high, steep, exposed cliffs occur along portions of the eastern shoresof the Naked Island group. Each is forested to its summit; the principal species of tree are Sitka spruce(Picea sitchensis), western hemlock (Tsuga heterophylla), and mountain hemlock (T.rnertensiana). All of these islands are partof the Chugach National Forest.
2 Naked I., about 30 !un southwest of the site where theT/V Exxon Valdez ran aground on Bligh Reef, was one of the first areas to be oiled (see Fig. 3, Kuletz 1994:35). Between 27 March and 2 April, 1989, portions of the eastern, northern, and northwestern shorelines were oiled. The prevailing winds moved most of the oil to the south, away from the island, but between7 and 9 April, southerly winds brought the oil into contact with the southern and western shorelines of Naked I. again. Neither Jackpot nor Fool Island were oiled.
Selection of Study Sites
We chose Naked I. as one of our principal study sites and as a base of operations. This island has been used as a base camp for several previous guillemot studies (Oakley and Kuletz 1979, 1993, Eldridge and Kuletz 1980, Kuletz 1981, 1983, Oakley 1981). We made numerous trips to other areas of PWS looking for additional colonies that might also serve as study sites. The two main criteria for determining the potential of a new guillemot colony as a study site were the number of breeding guillemots in that colony and the accessibility of the nest sites. We examined guillemot colonies in some of the fjords of northwestern PWS: Passage Canal, Blackstone Bay, the Barry Arm off Harriman Fjord, and the Yale Arm off College Fjord (Fig. 1). We also examined several island colonies: the Pleiades, Fool I., Jackpot I., Bligh I., and Hinchinbrook I. (Fig. 1). We investigated several colonies on Storey and Peak Islands of the Naked Island group, as well as many scattered around Naked I. itself.
Censusing: Population and Colony Attendance
Pigeon guillemot populations of Naked, Peak, Storey, Smith, and Little Smith Islands (the Naked Islandcomplex, Fig. 2) were censused by circumnavigating each island in a small boat at a distance of between 50 m and 100 m from the shore when the weather was good and the tides were near high. These censuses were conducted from 30 May to 1 June during the same timeof day (0400-1000 Alaska Daylight Time) and at the same timeof year that previous censuses of the this areawere made. Guillemots at Jackpot I. were censused near the high tide on 11 June, but at 1330 instead of in the morning. Guillemots around Fool I. were counted on 15 June at 0530, which corresponded to peak tide. Also, throughout the breeding season, butmostly during the chick-rearing period, counts of the maximum number of guillemots present at a particular colony were made at 15-minute intervals whenever thatcolony was being monitored from a boat or a blind. To examine colony attendance as a function of time of day, we made continuous 24-h watches at each of two guillemot colonies on Naked I. on 13-14 June and on 5-6 July. Observation shifts ranged from three to four hours.
3 Nest Sites and Monitoring
We marked 51 nests on Naked I., mostly at colonies along the western shoreline (Fig. 31, and 37 nests on Jackpot I. by painting a black number inside a white spot on nearby rocks. Some of the nests on Naked I. were known from previous studies, but most (65%)nests on Naked I. and all nests on Jackpot I. were found during the 1994 field season. We found some guillemot nests by searching potential nest sites in known colonies. Throughout the breeding season, but especially during the chick-rearing period, many new nests were opportunistically added to our sample as guillemots were discovered entering or leaving nest sites. Because of their inaccessibility or our inability to determine their contents, some of these nests were monitored only during feeding observations and were not used as part of our productivity sample. Nest sites were classified according to the type of habitat inwhich they occurred: tree root systems, rock crevices, or talus piles. On Naked I., we checked nests at approximately four-day intervals initially, and then about every two days during the fledging period. Because of the distance from our base camp to Jackpot I., we checked nests there at eight- to eleven-day intervals initially, and then at four- to five-day intervals or more frequently later in the season.
Banding and Morphometrics
Adults were caught in mist nets over the water, innoose mats on communal rocks, or by hand or net at the nest. Adults were banded on the left foot with a USFWS metal band (bottom) and a color plastic cohort band (top), and on the right foot with a unique combination of two color plastic bands. Chicks were banded on the right foot with a USFWS metal band (bottom) anda color plastic cohort band (top) andon the left foot with a unique combination of two color plastic bands. The 1994 cohort plastic band was yellow. We banded 19 adults and 61 chicks. All birds received a USFWS band; all adults and mostof the chicks also received the color bands. We measured all adults thatwe handled and all accessible chicks. We measured tarsus, culmen, and gape to the nearest tenthof a millimeter with vernier or dial calipers and maximum wing chord with a rule to the nearest millimeter. We weighed birds with Pesolam spring scales (0-100 g x 1 g, 0-500 g x 5 g, and 0-1 kg x 10 g) using the scale with the greatestprecision possible. Newly hatched chicks were marked on the right foot and on the down of their head with paint markers to distinguish between alpha (first-hatched) and beta(second- hatched) chicks until they were large enough tobe banded.
4 Nesting Chronology
Only nests that were discovered during the egg stage were used to construct the nesting chronology of guillemots at Naked and Jackpot Islands. For nests with two-egg clutches, eggs were assumed to have been laid on the same day and chicks to have hatched on the same day. If the laying date of the first egg of a two-egg clutch was known, the following day was used as the laying datefor both eggs. Laying dates were sometimes back-calculated from hatching dates using 31 days as the incubation period for both eggs. Oakley and Kuletz (1994) used 32 days and 30 days for the incubation periods of the first and second chicks, respectively. For determining fledging dates of chicks still present in the nest when we left the study area, achick was assumed to have fledged on the date of our last visit if the chick had been in the nest for 37 days or more. If the chick had not yet been in the nestfor 37 days, then it was assumed to have fledged on that date in the future equivalentto a 37-day nestling period, which is the mean reported by Oakley and Kuletz (1994)for Naked I.
Productivity
We estimated productivity by two different methods. During the incubation stage, a nest was considered to be active and included inour sample if it contained at least one egg and if an adult was seen in that nest at least once. If we knew two eggs had been laid in a nest but saw only one chick and no sign of the other egg, we assumed that both eggs hatched and one chick died. It seems unlikely that a predator entering an active nest would take only one egg and leave the other intact. Also, based on other guillemot studies (G.Divoky, personal communication; D.L. Hayes, personal observation), the proportion of two-egg nests in which only one egg hatches is fairly low. In the first method,we used only those nests thatwe followed from incubation through fledging. Productivity (chicks fledgedeggs laid) was defined as hatching success (eggs hatchedeggs laid) times fledging success (chicks fledgedeggs hatched). Thirty days is approximately the minimum time spent in the nest by guillemot chicks; the actual time is often much longer. For purposes of estimating fledging, however, any chick surviving in the nest for 30 days was assumed to have fledged. Because there must have been some nests we never found that had lost eggs or chicks, and consequently our estimates of productivity were probably biased high, we also estimated productivity using the methodsof Mayfield (1961, 1975). The Mayfield method eliminates some of this bias by recognizing that the number of nests lost will vary with the numberof nests in the sample and with the length of time for which the sample is monitored. The probability that an egg present at the startof incubation will produce a fledgling was calculated as the product of A (the probability an egg will survive a 31-day incubation period); B (the hatching success rate of those eggs present in the nest throughout the incubationperiod);
5 and C (the probability a chick will survive a 30-day nestling period). Values for B were calculated as in themethod described previously. Values for A and C were calculated from survival rates, which were based upon the "exposure" that each had in egg- or chick-days, respectively. When calculating exposure, if the number of eggs or chicks in a particular nest changed between visits, the change was assumed to have occurred at the midpoint of the interval, unless there was evidence to indicate otherwise, in which case the evidence was used to determine the date of the change. We used any nest for which we knew the contents on two or more dates, regardless of when the nest wasfound or how long we monitored it. This allowed us to increase our sample size and to make useof nests found later in the season.
Predation
Potential nest predators include the river otter(Lutra canadensis),mink (Mustela uison), northwestern crow (Coruus caurinus),common raven (C. corax), Steller's jay (Cyanocitta stelleri), and black-billed magpie (Pica pica). Bald eagles (Haliaeetus leucocephalus), peregrine falcons (Falco peregrinus), and other raptors might be predatory on adult and fledgling guillemots. We set Havahartm traps andchecked them daily in an attempt to capture and identify potential predators. We disguised three traps as burrows by covering all but the entrance withmoss, then baited them, first with eggs and later with fish. These were placed near guillemot nest sites in Cabin Bay that showed evidence of predation (an egg shell found in the open with a hole chewed in it but otherwise intact) and attempted predation (signsof scratching and diggingdirectly above an active guillemot nest). They were monitored from 19 June through 1 July, which approximately coincided with the latterportion of the incubation period. We reset the traps at a new location in Cabin Bay and monitored them from 8 July through 27 July, which marked the early stagesof the chick-rearing period. We did not disguise the traps at this new location and used various pieces of meat or fish as bait. If eggs disappeared from nests between visits,we assumed that predation was the cause. If chicks too young to fledge (i.e., younger than 30 days) disappeared from nests between visits,we assumed predation was the cause only if we were reasonably certain that no chick was still insome hidden corner of the nest. In some instances, where the nestcavity was too long or labyrinthine, it was not possible to make this determination. If after repeated visits to this type of nest, we never saw the chick(s) again, we listed the cause of failure as unknown.
Chick Growth and FledgingWeights
We calculated the growth rates of chicks as the change in weight (g/d) during the linear phaseof their growth, which is the period eight to 18 days after hatching (Koelink 1972). We measured few known-age chicks, so used wing length
6 as an index of developmental stage. Wing length can be measured easily and relatively consistently on chicks of all ages, especially after the feathersbegin to develop. Also, the wing grows continually from hatching to fledging and is less sensitive than body mass to short-term deficiencies in nutrition (Ricklefs and White 1975). A second-order polynomial was used to fit a curve to the data, where In body mass was plotted as a function of In wing length. The comparison between the growth rates of chicks on the two islands is confounded by the availability of different numbers of measurements on different chicks and different numbers of chicks in different nests. Thus, there is a complicated structure of interdependence among observations on each island. If this were not so, and observations from each chick were independent from those of other chicks on that island, then itwould be possible to compare curves relating In body mass to In wing length by standard regression methods. Therefore, a randomization-based approach was used (Manly 1991). Preliminary analysis indicated that the relationship between body mass (B) andwing length (W) is well approximated on each island by an equation of the form ln(B) = Bo + B, * ln(W) + B, * ln(W)'. (1) Assuming this to be so, a measure of the extent towhich the data arefitted better by separate equations for each island than by a single equation for both islands is the reduction in thecoefficient of multiple determination (R'), obtained when separate equations are fitted rather thana single equation. A test for whether this is a reduction is achieved by determining whether theobserved reduction is significantly large in comparison to the distribution obtained for the same statistic by randomly reallocating nests (with all their associated measurements on one or two chicks) between the two islands, while keeping the number of nests on each island equal to theobserved number. Fledging weight was assumed to be the last recorded weight of a chick that was measured within one week of its departure from the nest. Chicks still in the nest when we left the study area werenot included in the calculation of fledging weights.
Chick Provisioning and Diet
Either from blinds or from boats we observed adult guillemots bringingfood items to their chicks throughout the chick-rearing period. Feeding watches ranged from 1.25 h to 24 h for a total of 388.75 h at Naked I. and 82.5 h at Jackpot I. Usually just one observer was in theblind at a time; shifts ranged upto six hours). Binoculars and spotting scopes were used to identify prey items in the bills of guillemots to the lowest possible taxon or "type" of prey. When time and visibility permitted, we also estimated the lengthof the prey item as a multiple of the guillemot's bill to the nearest half bill length. We recorded the time of each delivery and the numberof the nest to which the prey was delivered, as well as how long the adult first remained on the water with thefish before delivering it.
7 We also recorded instances of attempts at kleptoparasitism or predation by gulls and raptors. To test whether deliveries were distributed more or less evenly throughout the daylight hours, the day was divided into three approximately equalperiods: early (0600-1100), mid-day (1100-1700), and late (1700-2200). Although a few deliveries occurred very early and very late, when itwas too dark to make reliable observations, the period used for analysis was truncated at both ends in accordance with the times listed above. Using a Chi-square goodness-of-fit test, the actual number of deliveries observed during each of the three periods was compared to the expected number of deliveries in those periods if they had been distributed evenly throughout the day.
Identification and Samplingof Fish Types
We occasionally sampled waters (< ca. 15 m) around Naked I. with fish traps set on the bottom or occasionally above it to obtain specimens of fish to aid us in identifylng those in the bills of guillemots. We measured wet weight and total length of all fish. Unless used as an identification sample, each fish was released after being weighed and measured. Occasionally, we retrieved fish of species not caught in the trapsthat we found in or near guillemot nests and used these as identification samples.
Data Analysis
Comparisons between Naked and Jackpot Islands or between years were made with two-tailed t-tests, 2 x 2 contingency tables analyzed with a G-test and corrected for continuity, Chi-square tests, ratio estimation with Z statistics, and randomization techniques. The level of significance was set at a = 0.05. All means are reported as the mean plus or minus one standard deviation.
RESULTS
Censusing: Population and Colony Attendance
In 1994, 1,262 pigeon guillemots were counted around the shorelinesof the Naked Island complex during the early-season census (30May - 1 June, Table 1). On 11 June, 34 guillemots were counted around Jackpot I. The maximum number counted, however, was 74 at 0530 on 6 August. This latter count was conducted from an anchored boat during feeding watches and thereforeonly includes birds seen from that vantage point. The birds were in several large rafts, gradually moving closer to the island as daylight increased. Thus, it is possible that these rafts included most of the Jackpot I. population. On 15 June,74 birds were counted around Fool I.
8 During the incubation period (Fig. 4) and early hatchingperiod (Fig. 51, maximum counts of pigeon guillemots usually occurred in the early morning hours, shortly after first light. The birds were first detected in rafts a considerable distance from shore, then graduallymoved closer to the colony. Time of day appeared to be more important than the tidalcycle in affecting colony attendance. Secondary peaks of guillemots later in the day often corresponded to high tide. Although no continuous 24-h observations were made in August, numerous counts made during extended feeding watches suggest that the tidal cycle had much less effect on colony attendance of guillemots while they were provisioning their young. The maximum number of guillemots counted at the colonies we studied around Naked I. (Fig. 3) was as follows: Igloo I(37 on 26 July), Igloo I1 (14 on 2 August), Nomad (27 on 8 August), Row (20 on 10 July), Thumb (9on 30 May), Hook I (17 on 23 July), and Hook I1 (11 on 24 July). The numbers of active guillemot nests at five study colonies on the western sideof Naked I. are compared with similar data from previous years in Table 2. Since the last count in 1990, the numberdeclined at three colonies and increased at two, although the number at every colony has declined since the 1978 count. The combined number of active nests at all five colonies in 1994 was only about half the numberfound in 1978 and 1979. Of the 51 marked nest siteson Naked I., 15 (29%) were among tree roots, 28 (55%) were in rock crevices, and eight (16%) were in the talus. All but one of the 37 marked nest sites on Jackpot I. were among tree roots and the remaining one was in a rock crevice. There were few suitable rock crevices on Jackpot I. and no talus piles.
Nesting Chronology
Nesting chronology was similar at both islands (Naked I., Fig. 6.; Jackpot I., Fig. 7). The median laying date was4 June at Naked I. (range = 19 May - 17 June) and 3 June at Jackpot I. (range = 24 May - 13 June). The median hatching date was 3 July at both Naked I. (range = 19 June - 18 July) and Jackpot I. (range = 24 June - 14 July). The median fledging date was 14 August at Naked I. (range = 2 August - 24 August) and 9 August at Jackpot I. (range = 30 July - 21 August). However, out of this sample, two chicks from each island were still present in the nest on our last dayof observation (Naked I., 22 August; Jackpot I., 18 August). The mean number of days that chicks spent in the nest wasnot significantly different between the two islands (t = 1.922, df = 43, P > 0.05). Chicks were in the nest an average of 39.3 5.5 d on Naked'I. (n = 18, range = 30-48 d) and 36.6 * 3.8 d on Jackpot I. (n = 27, range = 30-47 d).
9 Productivity
The mean clutch size was 1.70 f 0.47 (n = 23) on Naked I. and 1.92 f 0.28 (n = 24) on Jackpot I. The difference between clutch sizes on the two islands was not significant (G = 2.480, df = 1, P > 0.10). Of a total of 39 eggs (23 clutches; 16 with 2 eggs, 7 with 1 egg) on Naked I., 35 hatched, two were abandoned, and two were probably taken by predators. Of a total of 46 eggs (24 clutches; 22 with 2 eggs, 2 with 1 egg) on Jackpot I., 37 hatched and nine were abandoned. Of a total of 53 chicks (32 broods; 21 with 2 chicks, 11 with 1 chick) on Naked I. that were monitored at some stage of the breeding cycle, 30 chicks fledged, eight were found dead in the nest, 11 were probably taken by predators, and the fateof four others is unknown. Of a total of 45 chicks (23 broods; 22 with 2 chicks, 1 with 1 chick) similarly monitored on Jackpot I., 35 chicks fledged, three were found dead in the nest, and the fateof seven others is unknown. The difference between brood sizes was not tested for significance because we were not sure of the original number of chicks in some of the "one-chick" nests found after the egg stage. Hatching and fledging success and productivity, as determined by the "traditional" method, were basedon 23 clutches from 22 nests (one pair relaid after their nest was depredated during theegg stage) at Naked I. and 24 nests on Jackpot I. Hatching success was 0.90 (n = 39) at Naked I. and 0.80 (n = 46) at Jackpot I.; it was not significantly different between islands (Z = 0.90, P = 0.1841). Fledging success was 0.51 (n = 35) at Naked I. and 0.76 (n = 37) at JackpotI.; it was significantly different between islands (Z = 2.13, P = 0.0166). Overall productivity (i.e., chicks fledgedeggs laid) was 0.46 at Naked I. and 0.61 at Jackpot I. The Mayfield method yielded similar productivity values of 0.42 (n = 36) at Naked I. and 0.61 (n= 29) at Jackpot I. Fourteen of 23 nests (61%)at Naked I. produced at least one fledgling compared to 18 of 24 nests (75%) at JackpotI. This difference between the proportion of successful nests was not significant (G = 0.528, df = 1, P > 0.25) between the two islands.
Predation
Of the 35 nests on Naked I. that we monitored for productivity, two nests were depredated during theegg stage and eight nests were depredated during the chick stage. Although potential predators were seen on both islands, none were actually seen taking eggs or chicks. There was no evidence of predation on Jackpot I., but we found evidence suggesting mammalian predation on Naked I. in the form of guillemot carcasses with the heads chewed off, signs of scratching and digging at nest entrances, and mammalian hair€ound near a depredated nest. Guillemots often reacted to the presence of bald eagles around the colonies by flushing from the rocks or nearshore waters and settling farther offshore. Gulls and raptors were occasionally seen chasing guillemots with food, but in almost all instances, this behavior appeared to be kleptoparasitic rather than predatory.
10 Black-billed magpies were often seen near guillemot nest sites and were sometimes seen entering the nests. The shells of several apparently depredated guillemot eggs were also found, but it wasnot possible to identify the predator from this evidence.
Chick Growth and Fledging Weights
The mean growth rateof chicks during the linear phaseof their growth was 15.7 * 6.6 g/d (n = 10, range = 5.0-29.0 g/d) at Naked I. and 20.3 k 3.5 g/d (n = 6, range = 15.0-23.5 g/d) at Jackpot I. The difference was not significant (t = 1.570, df = 14, P > 0.10). The relationship between In body mass and In wing length of chicks on Naked and Jackpot Islands is compared in Figure 8. Fitting a common curve of the form of equation (1)to the results for both islands gives R2 = 0.9284. Fitting separate curves gives R2 = 0.9352. The increase in R2 obtained by fitting separate curves is therefore 0.0068. To test whether this increase is significantly large, the randomization distribution of the increase inR2 was approximated by the observed value plus 999 values obtainedby randomly reallocating the available data set of 41 nests to a set of 24 nests on Naked Island and 17 nestson Jackpot Island. Only 12 of the 1000 values were equal to or greater than 0.0068. Therefore, the increase of 0.0068 is significantly large (P= 0.012), clearly indicating a difference in the body mass - wing length relationships for the two islands (Manly 1991). The mean fledging weight of chicks was 453 k 55 g (n = 17, range = 357-525 g) at Naked I. and 508 f 39 g (n = 17, range = 440-585 g) at Jackpot I. This difference was significant (t = 3.410, df = 32, P e 0.002).
Chick Provisioning and Diet
Collectively, guillemots delivered fish to their chicks throughout the daylight hours at Naked I. (Fig. 9) and at Jackpot I. (Fig. 10). Neither distribution was significantly differentfrom a theoretical even distribution of deliveries made throughout the day (NakedI., x2 = 3.709, df = 2, P > 0.10; Jackpot I., x2 = 0.936, df = 2, P > 0.50). Feeding rates varied considerably among nests. At any particular nest, there wereperiods of several hours in which no deliveries were made; also adultssometimes stopped delivering shortly after mid- day (Naked I., Figs. 11 and 12; Jackpot I., Fig. 13). Pigeon guillemots at Naked I. sometimes foraged directly in front of their colony in water less than 15 m deep, but usually foraged in nearby bays or on the broad, shallow-water (e 25 m deep) shelves surrounding Naked I. (Fig. 14). Guillemots were never seen foraging in the immediatevicinity of Jackpot I., but instead flew toward shallower waters near the mouthsof Jackpot Bay or Icy Bay (each ca. 2-4 km distant, Fig. 15), presumably foraging there. Guillemots carrying fish were observed returning to the colony at Jackpot I. from the general direction of either of these two bays.
11 The diet of pigeon guillemot chicks at the two islands was different with respect to types of fish. At Jackpot I., herring or smelt accounted for at least 32.3% of the prey items fed to chicks, whereas at Naked I., these types of fish accounted for only 1.2% of the chick diet (Fig. 16). Gadids were prevalent in the diet of chicks at both islands (Naked I., 29.5%; Jackpot I., 18.5%). Sand lance accounted for at least 7.9% of prey items delivered at Naked I., but were virtually absent (0.8%) from the chicks' diet at Jackpot I. Because of the relatively large proportion of fish that could not be identified (Naked I., 25.4%; Jackpot I., 24.2%), the values reported above represent minimum percentage contributionsof those types of fish to the total delivered.
Fish Types Caught in Traps
Shrimp (mostly Pandalus danae and Eualus gaimardii) were the most frequently taken animal in the fish traps, but werenot counted because they were never seen being delivered to guillemot nests in 1994. Of 106 fish caught in the traps, the relative proportions of each type were as follows: 40 arctic shannies (Stichaeus punctatus),25 gadids (presumably mostly Pacific tomcod Microgadus proximus, possibly also Pacific cod Gadus macrocephalus, and walleye pollock Theragra chalcogramma), 19 Lumpenus spp. (probably mostly snake pricklebacks L. sagitta), nine sculpins (mostly tidepool sculpins Oligocottus maculosus, also one Nautichthys sp.), eight crescent gunnels (Pholis laeta),three masked greenlings (Hexagrammos octogrammus),one juvenile lingcod (Ophiodon elongatus),and one warbonnet (Chirolophis sp.).
DISCUSSION
Censusing: Population and Colony Attendance
Early season counts of pigeon guillemots in the Naked Islandcomplex suggest that their population has decreased considerably from 1978 and 1979 (Table 1). These censuses, which were conducted at the breeding colonies during roughly the same phase of the breeding cycle, at approximately the same time of day, and under similar weather and tidalconditions, probably are better indicators of trends in guillemot numbers than are counts made during other types of surveys in PWS. Observed colony attendance patterns of guillemots at Naked I. (Figs. 4 and 5) and Jackpot I. indicate that the timeof day is extremely important when planningguillemot censuses. A morning peak in colony attendance seems to hold true for pigeon and black guillemots throughout most of their range (Drent 1965, Cairns 1979, Vermeer et al. 1989). Vermeer et al. (1989) reported that the optimal time to determine the population of nesting guillemots was at high tide in the morning. They found colony attendance to be correlated most strongly with tide height, but also with timeof day and tide direction,at three colonies in Skidegate Inlet, Queen Charlotte Islands. However, at Mandarte
12 Island, British Columbia, time of day was theonly factor influencing colony attendance. They suggested that the maximum tidal range was responsible for the observed differences in colony attendance. The range at Skidegate Inlet is 7.8 m, but only 3.8 m at Mandarte Island; it is 5.8 m in the Naked Island area. Another indication of a marked decline is the total numberof active nests at five previously monitored colonies on Naked I., which in 1994 was only about half of what it was in 1978 and 1979 (Table 2). Other surveys such as those of Dwyer et al. (ND;ca.15,700 in 1972-19731, those of Klosiewski and Laing (1994; ca. 3,000-6,600 in 1989-1991), and most recently by Agler et al. (1994; ca. 4,000 in 1993) also suggest a general decline in the numbers of pigeon guillemots in PWS. Sanger and Cody (1994) surveyed 98% of the PWS shoreline between May and June of 1993 and counted 3,028 guillemots. Because the primary objective of their survey was to locate pigeon guillemot colonies, their methods were differentfrom the previous censuses conducted around the Naked Island complex, and the lower count for 1993 in this area most likely reflects these differences in methods. A comparison of the counts by Sanger and Cody (1994) with other recent counts (see Table 1) made before and after suggests that their estimates mightbe about 35% low.
Productivity
The ideal and most straightforward method of calculating productivity is from a sample of known nests that are followed from before egg-laying through fledging. Use of this "traditional" or cohort method was not an option during the 1994 field season. First, in most instances, we did not know about a nest site until we discovered eggs or chicks, because guillemots build no nest and use only a rudimentary nest scrape inwhich to lay their eggs. Second, we found most nests either well into the incubation period or during the chick-rearing period, when parents were observed delivering food to their chicks. If all nests that were found later in the breeding season are included in the sample,one runs the risk of biasing the results toward successful nesting attempts. This is because those nests that failed without ever having beenfound are not accounted for in calculations of productivity. As a compromise, we included in our sample those nests whose contents were monitored from incubation through fledging. The problem with this method is that we are restricted to small sample sizes (Naked I., 23 clutches from 22 nests; Jackpot I., 24 nests). The cryptic nature of guillemot nests makes it likely that in most studies (especially during the first year),a large proportion of the nests monitored will be those that are discovered later in the breeding season. The Mayfield method allowed us to use those nests found later in thebreeding season and thus increase our sample sizes (Naked I., 36 clutches from 35 nests; Jackpot I., 29 nests). The Mayfield method provides a useful means of comparing data between studies when appropriately sized cohorts cannot be followed throughout the breeding season.
13 Oakley and Kuletz (1994) included several tables in their report, each of which relates to some measure of pigeon guillemot productivity. These tables have been reproduced (see Appendix, Tables Al-A5) with only minor modifications in this report, and updated with the appropriate valuesfor 1994. Although the original tables contained information from Naked I. only, data from Jackpot I. have been included for comparative purposes.
Predation
From our productivity sample, nests of each type were depredated in 1994: tree roots (4 of 10, or 40%),rock crevices (4 of 19, or 21%),and talus (2 of 6, or 33%). In 1978, of 146 known nest sites on Naked I., 40% of the nests were among tree roots, 36% in rock crevices, and 24% in talus (Oakley 1981). The high proportion of nests in rock crevices (55%,usually on cliff faces) on Naked I. in 1994 suggests that guillemots might be selecting this type of nest site tobe less accessible to predators. The difference in the proportion of nests of each type between 1978 and 1994 was not significant (x2= 5.455, df = 2, P > 0.05). However, the number of rock crevice nests in 1994 was higher than the expected and this component made the largest contribution to the calculated Chi-square value. Not all of these nests were accessible, so we could not determine the fate of eggs or chicks in some. Jackpot I. has little shoreline (ca.1 km)and low cliffs (typically < 3-4 m high) with few suitable crevices and no talus piles. Therefore, almost all guillemots breeding on the island use tree roots as nest sites. Although all or most of the tree root nests on Jackpot I. were seemingly accessible to mammalian predators, we found no evidence of predation there. In Shetland and Orkney, the use of available breeding habitat by black guillemots was influenced by the distribution of mammalian ground predators (Ewins and Tasker1985). Nest sites accessible to ground predators were usedon islands that were free of brown rats (Rattus noruegicus) and ermine (Mustela erminea),but on islands where these mammals were present, guillemots bred only in higher cliffs. Oakley and Kuletz (1994) noted that the primary difference in productivity of pigeon guillemots on Naked I. that they observed following the oil spill was lowered nesting success, which was the resultof nest predation during the chick stage. On Naked I. in 1994, we found carcasses of guillemot chicks with the heads chewed off, suggesting that some kind of mustelid is likely responsible for the predation. The most likely predator is the river otter. These animals were seen frequently around Naked I. in the vicinity of guillemot colonies. Ewins (1985) reported that on the island of Mousa in Shetland, otters(Lutra lutra)killed both chicks and incubating adults, and that decapitated carcasses were a sure sign of these predators. Ewins also noted that there were few nests inaccessible to them. Likewise, many of the nest sites on Naked I. are probably accessible to otters, including some of those in rock crevices. Few, if any, nest sites would be inaccessible to the more agile mink. We never saw mink in 1994. Although mink were common on Naked I. during the
14 summer after theoil spill, no mink were seen in 1991 and only scant evidence of their presence was found in 1992 despite some trapping effort (J. Faro, personal communication). River otters and mink typically forage in the intertidal zone. A study that compared the diet of river otters intwo areas of PWS before and after the spill showed that there were significant declines in species richness and diversity (mostly bony fish and molluscs) in otter diets on the oiled area compared to the unoiled area (Bowyer et al. 1994). Another study, also conducted in PWS, clearly indicated that oil contamination was affecting the health of river otters up to two years after the spill (Duffy et al. 1993, 1994). Contamination of the normal intertidal food supply of river otters and mink might have ultimately causedsome of these predators to switch to other types of prey, including guillemotchicks. Whatever mammalian predator is responsible for killing chicks on Naked I., the increased predation pressure there might have caused breeding guillemots to move elsewhere. It is possible that guillemots in PWS are emigrating from some colonies on the mainland and large islands like NakedI. to smaller ones like Jackpot I., where ground predators have not become established. Emigration of black guillemots from colonies in Sweden and Iceland have been attributed to predation by mink (Asbirk 1978, Petersen 1979). These kinds of movement would not necessarily show up in Sound-wide surveys such as those of Agler et al. (19941, but rather incensuses designed to count guillemots at their colonies during the breeding or prebreeding season. All black guillemot colonies in Scotland were surveyed in the mid-1980's using standardized methods(Le., early morning counts in the prebreeding period; M.L. Tasker, personal communication). In several of the areas, more recent counts have provided useful indices of change. The principal factor controlling local distributions in Scotland appears tobe introduced mammals. Avian predation also probably accounts for some of the nesting failureson Naked I. Of the corvids, magpies are the likeliest predator. They were seen on several occasions either entering guillemot nests or repeatedly following adult guillemots to the nest entrance. During a previous field season, a magpie was seen dragging a guillemot chick from its nest (K. Kuletz, personal communication). Crows are apparently awareof the locations of marked guillemot nests and were seen in 1994 and in previous years "systematically"moving from one marked nest to another. Other studies indicate that crows are a major source of egg predation and sometimes take young chicks as well (Emms and Verbeek 1989, Ewins 1989). Adults, and especially fledglings, are probably sometimes taken by large raptors. Bald eagles are known predators of adult guillemots in British Columbia (Vermeer et al. 1989 as cited in Ewins et al. 1993). Beaks of guillemots were found beneath an eagle's neston Naked I. during a previous'study (K. Kuletz, personal communication). We often witnessed a change in the guillemots' behavior when an eagle flew into the area. The guillemots' reactions to the presence of bald eagles (e.g., flushing, moving farther offshore, alarm calling, and diving) suggest that they perceive this potential predator as a real threat. However, any interactions between an eagle and a single guillemot were assumed
15 to be kleptoparasitic because we only observed these when a guillemot had a fish in its bill.
Other Causes of Nesting Failure
Only two eggs from a single nest were abandonedon Naked I., but nine eggs from five nests were abandoned on Jackpot I. The abandoned nest on Naked I. was in a crevice that had little cover. Abandoned nests on Jackpot I. were typically deep cavities in expansive tree root systems. An adult was flushed twice from the nest on Naked I. and twice from one of the nests on Jackpot I. It is possible that our repeated visitsto these nests was the causeof this abandonment. Several investigators at other guillemot colonies have observed reduced productivity apparently associated with human disturbance (Bergman 1971, Cairns 1980, Vermeer et al. 1993). The fact that the rateof abandonment on Jackpot I. was higher, although nests were visited more frequently on Naked I., might be explained by the lack of previous research on Jackpot I. On Naked I., some of the same colonies have been studied for years and the guillemots there might have habituated to some extent to the presenceof biologists walking through their breeding habitat. One chick on Naked I., and two chicks on Jackpot I., died within their first week of life. The cause of death in each instance was unknown. Other studies have also reported deaths among young chicks (Preston 1968 and Kaftanovski 1951 as cited therein; Divoky et al. 1974, Kuletz 1983, D.L. Hayes, personal observation). Preston (1968) suggested the cause might be related to prolonged chilling or failure to become homoiothermic. On Naked I. in 1994, two chicks apparently died from exposure after heavy rains and one chick died after its wing had been pinned down by a fallen rock.
Chick Growth and Fledging Weights
Our estimates of growth rates during the linear phaseof growth (Naked I., 15.7 g/d; Jackpot I., 20.3 g/d) were similar to those of Oakley and Kuletz (1994) at Naked I. (range = 16.6-23.8 g/d for all years combined, Table 3), as were our estimates of fledging weights (Table 4). Growth rates were also similar to those reported by Koelink (1972) at Mandarte Island (15.9 g/d) and Ainley and Boekelheide (1990) on the Farallon Islands (16.5 g/d). In the absence of known- age chicks, we used our plots of In body mass vs. In wing length as a condition index to the developmental stage of the chicks. Manipulation of food intake of alcid chicks by several researchers has shown that the rateof growth of wing feathers is unaffected by nutrition (Gaston 1985 andreferences therein). Randomization techniques showed that the two data sets graphically compared in Figure 8 were significantly different. On average, chicks on Jackpot I. also fledged at greater weights and spent less timein the nest than those on Naked I.
16 Peakall et al. (1982, 1983 as cited in Oakley and Kuletz 1994) have shown that ingestion of a small amount of oil by waterbirds can produce a variety of sublethal, physiological effects that might affect their fitness. Black guillemot chicks dosed with crude oil and raised by their parents in the nestgrew significantly more slowly than controls, and also showed signs of impaired osmoregulatory function (Peakall et al. 1980). Although it is unlikely that direct ingestion of oil or external oiling is affecting the birds five years after the spill, indirect effects of the oil might be important. Hemosiderosis has been observed in demersal fish collected from oiled eel grass beds in Herring Bay, Knight Island, PWS (S. Jewett, personal communication). Hemosiderin is the result of excessive destruction of erythrocytes and collects in the liver and spleen. Polycyclic aromatic hydrocarbons (PAHs) found in crude oil are known to cause the lysis of red blood cells. Of a sample of 10 pricklebacks (Anoplarchus spp.) and 10 crescent gunnels (Pholis laeta) collected from each of two areas of the bay (oiled and unoiled), all specimens from the oiled beds showed signs of hemosiderosis and none in the control beds showed any signs of this disorder. Many of the fish from the oiled beds were in poor condition, as judged by lipid and glycogen stores. Long-term exposure of marine fish to crudeoil results in lowering of lipid reserves (Thomas et al. 1980 and Dey et al. 1983, both cited in Duffy et al. 1993). If guillemots are regularly feeding their offspring fish from oiled areas, the chicks could be accumulating PAHs in concentrations that might cause sublethal toxic effects. Either reduced energy content of food fish or accumulation of PAHs could adversely affect chick metabolism and growth. Differences in diet might also explain thedifferences in growth rates between the two populations. The energy density of lipid is about twice that of protein or carbohydrate. The lipid content of herring, capelin, and sand lance is typically high relative to thatof gadids, which tend to be poor in lipids (D. Roby, personal communication). Also, research on adult kittiwakes and murres by Brekke and Gabrielsen (1994) suggests that assimilation efficiency is directly proportional to the fat content of the prey. This same relationship might hold for some seabird chicks as well. The relative proportions of these types of fish delivered to guillemot chicks (especially the large component of herring or smelt at Jackpot I.) suggest thatchicks raised on Jackpot I. are, on average, receiving meals of higher energy content than those chicks raised on Naked I. The mean fledging weight of guillemot chicks raised on Naked I. (453 g) was 95% of the mean adult weightof 477 * 38 g (range = 420-569 g, n = 19, SE = 8.8), while the mean fledging weight of chicks raised on Jackpot I. (508 g) was 106% of this value. Supra-adult weights of chicks are not common among the alcids (Sealy 1973). Other studies have shown that pigeon guillemots typically weigh less than adults when they fledge and that their peak weight is reached at fledging. On Mandarte Island, the meanfledging weight was 411 g, or 91% of the mean adult weight (Drent 1965); in Puget Sound, the meanfledging weight was 438 g, or 92% of the mean adult weight (Thoresen and Booth 1958). The fact that the mean fledging weight of Jackpot I. chicks was greater than the mean adultweight might
17 be explained by the fact that all the adults measured werefrom Naked I. and the mean adult weight of the Jackpot I. populationcould be greater, or that sample sizes were too small and thus had too much variance. However, Eldridge and Kuletz (1980) reported a mean adult weight of 504 & 25 g (n = 42, SE = 3.8) for Naked I. pigeon guillemots that they measured in 1979; mean fledging weights were 506 2 38 g (n = 18 chicks measured within four days of fledging) and 503 * 43 g (n = 8 chicks measured within 24 hours of fledging). When making these comparisons, it is also important to remember that the massof chicks can vary considerably (up to 30-40 g/d) during the finalportion of the nestling period (Emms and Verbeek 1991). It seems reasonable to assume that inspecies such as the pigeon guillemot, where the chicks are independent upon fledging, fledging weight would be important to survival (i.e., heavier chicks with larger fat storeswould have an added buffer while learning to forage efficiently, especially if food is scarce or bad weather prevents them from foraging). There are no conclusive data indicating that fledging weight in auks influences their postfledging survival (Harris and Rotherby 1985). Other studies of species whose chicks are independent at fledging (e.g., Manx shearwater, Perrins et al. 1973; cape gannet, Jarvis 1974 as cited in Harris and Rotherby 1985) have shown that there is a significant positive correlation between weight at fledging and postfledging survival (but see Schreiber 1994).
Chick Provisioning
Members of the genus Cepphus typically lay two eggs. Most other alcids lay only a single egg, but the near-shore foraging habitsof guillemots probably account for their ability to raise two chicks. Mehlum et al. (1993)maintain that long-distance foraging by black guillemots, which typically raise two-chick broods and have a high wing loading relative to most other seabirds, is too energetically demanding and might exceed their maximum sustainable working level. Koelink (1972) argues this same pointfor chick-rearing pigeon guillemots. In his study, although artificial broods of three were successfully raised to fledging, there was a proportional decrease in the amountof food delivered per chick throughout the nestling period. In black guillemots also, artificial triplets have been successfully raised but with differing results regarding fledging weights. In Denmark, the mean fledging weight of triplets was higher than thatof chicks from normal broods (Asbirk 1979 as cited in Harris and Birkhead 1985). In Iceland, triplets fledged at lower mean weights than chicks from normal broods (Petersen 1981 as cited in Harris and Birkhead1985). Our measured rates of food deliveries to individual nests (range = 0.67- 1.39hesth; Naked I., Figs. 11 and 12; Jackpot I., Fig. 13) are comparable to those of other studies of Cepphus guillemots (Thoresen and Booth 1958, Bergman 1971, Asbirk 1979 as cited in Harris and Birkhead 1985, Cairns 1981, 1987, Kuletz 1983). Unfortunately, we did not always know the number of chicks inside each
18 nest. Without this information, and without a knowledge of the weight of each prey item delivered, a comparison of provisioning rates (i.e., g/h/chick) is impossible. Furthermore, fish varyconsiderably in their composition of lipids, proteins, and carbohydrates. Fish higher in lipids have a higher energy content, which can be particulary relevant to the reproductive success of the seabirds feeding upon them. Also, the lipid content even within a single species of fish can vary widely with season, sex, reproductive status, and age class(D. Roby, personal communication). It is almost impossible to accurately estimate the weight of prey items delivered to chicks noninvasively. Measuring the actual energy contentof the prey cannot be done by noninvasive means; prey must be intercepted and analyzed in the laboratory. Obviously, this cannot be done repeatedly at the same nest without affecting the food intake of the chicks involved.
Foraging
The maximum diving depth of black guillemots is about 50 m (Piatt and Nettleship 1985). Assuming that the pigeon guillemot has similar diving capabilities, it is restricted to waters no deeper than this whenfeeding on benthic prey items. The pigeon guillemots breeding on Naked I. generally forage around the island, usually within about600 m of the shore and in water shallower than 25 m (Kuletz 1983). There is a broad, shallow-water shelf surrounding Naked I. and the neighboring islands (Fig. 141, which allows guillemots to forage nearby. On Jackpot I., thereis very little shallow water immediately around the island (Fig. E), and thus guillemots breeding there fly greater distances to obtainfood for their chicks. These birds apparently find it necessary to fly several kilometers to Jackpot Bay or Icy Bay to find food for their chicks instead of trying to forage around Jackpot I. However, the extra amount of energy expended by adults feeding chicks on Jackpot I. seems to be offset by the increased growth rates, fledging weights, and productivity measured at this colony relative to colonies on Naked I.
Chick Diet
The most obvious difference in the diet of chicks between Naked and Jackpot Islands in 1994 was in the proportion of herring or smelt (Fig.16), accounting for approximately one third of all prey items delivered to Jackpot I. chicks, but only slightly more than one percent of those delivered to Naked I. chicks. Interestingly, herring or smelt were not noted in the diet of chicks on Naked I. in 1979 or 1980, then accounted for about 16% and'23% of the diet in 1981 and 1989, respectively, and only about two percent in 1990 (Oakley and Kuletz 1994). In 1994, sand lance accounted for less than one percent of all prey items delivered to chicks on Jackpot I., and about eight percent on Naked 1. Data from before the oil spill show that sand lance hadbeen the single most important prey item in the diet of chicks at Naked I. (55%in 1979, 35% in 1980, and 23% in
19 1981; Kuletz 1983). After the spill, in the two years when data are available, sand lance accounted for about 14% and eight percent of the chick diet (Oakley and Kuletz 1994). In 1994, gadids, which were relatively unimportant in the chick diet at Naked I. until after the spill, were the most frequently delivered type of prey at Naked I., and second only to herring or smelt at Jackpot I. The proportion of herring or smelt in the diet of chicks might be related to the ephemeral nature of schools of this type of fish and their presence within the foraging range of guillemots. Their capture might occur only conicidentally when behavioral factors (e.g., spawning) or oceanographic factors (e.g., currents, upwelling) bring these prey into shallower nearshore waters. However, the relative increase in the proportion of gadids, presumably caught by the guillemots on or near thebottom, could indicate a pronounced shift in the ecosystem. The fact that gadids did not show up in fish traps in appreciable numbers (Kuletz1983, Oakley and Kuletz 1994) until 1994 lends support to this hypothesis.
CONCLUSIONS
We found important differences between the pigeon guillemot populations of Naked and Jackpot Islands. On Naked I., we noticed increased predation pressure and a marked change in the diet of chicks compared to the late 1970's. Also, the numbers of guillemots in the Naked Islandcomplex are about half of what they were in the late 1970's. Productivity was lower on Naked I., primarily asa result of predation during the chick stage. The most likely predators are mustelids, especially river otters. Although we saw the same potential predator species on both islands and almost all of the nests on Jackpot I. appeared to be accessible to ground predators, we found no evidence of predation on that island. Egg abandonment was the main reason for nesting failures on Jackpot I. Our investigations might have caused the incubating adults to desert. The diet of chicks was substantially different between the two islands with respect to the proportion of schooling fish. Herring or smelt accounted for about one third of the chick diet on Jackpot I., but only about one percent on Naked I. The guillemots breeding at Jackpot I. foraged farther from their colonies than those breeding at Naked I. The extra energy expended by the Jackpot I. adults when foraging at greater distances seemsto be offset by increased growth rates of chicks and greater fledging weights. It is conceivable that residual oil contamination has caused river otters or other mustelids on Naked I. to switch to another food source. It is also possible that oil contamination has had sublethaltoxic effects on both demersal fish and, at the next trophic level, on the guillemots that feed on them. In addition to accumulating polycyclic aromatic hydrocarbons in their tissues, stressed fish are likely to have lowered glycogen and lipid stores, and thus energy content. These trophic relationships certainly merit further study. Sand lance were the singlemost important species in the diet of pigeon guillemot chicks on Naked I. in the late 1970's, but they accounted for only about
20 eight percent of the chick diet in 1994. Likewise, gadids were previously unimportant in thechick diet on Naked I., but in 1994 accounted for about 30% of the prey items delivered to chicks. The apparent decline in the availability of sand lance and changes in the relativeproportions of benthic and schooling fish in the diet of guillemot chicks might represent a key change in the ecosystem that is affecting other marine birds and mammals inPWS. Future work on pigeon guillemots in PWS should focus on 1) determining what animals areresponsible for the increased levels of predation at the nest site on Naked I., 2) marking chicks and breeding adults for estimating recruitment and adult survival, 3) censusing designated colonies on a regular basis using standardized methods, and 4) diet composition and energy content of prey items as they relate to growth and productivity.
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28 Table 1. Counts of pigeon guillemots during June censuses at Naked, Peak, Storey, Smith, and Little SmithIslands, Prince William Sound, Alaska, before and afterT/V the Exxon Valdez oil spill. Censuses conducted between 3 and 6 June unless otherwise noted. Dashes indicateno surveys were conducted.
Little NakedStoreySmithSmithPeak Yea r IslandYearIsland Island Island Island Total Before spill 1978 1115 392 1965 9472 175 1 979 1226 495 150 301 58 2230 58 301 150 495 1226 1979 1 980 891 __1980 ______
After spill 19 89" 615 193 73 193 615 1989" ______19 90 729 293 102 124 102 293 729 1990 31 1279 19 91 755 293 102 293 755 1991 1261 35 76 1 992 586 230 87 230 586 1992 100 23 1025 1993' 242 385 828 9432 75 1994' 298 739 81 1211262 23 "Census conducted on 13-14June. 'From Sanger and Cody 1994 (censuses in MayJune or 1993) 'In 1994 Naked Island census was done onMay, 30 Storey and Peak Islands on 31May, and Smith and Little Smith Islands on June1. Note: Data from all years except 1993 and 1994 from 1 Table (Oakley and Kuletz 1994).
29 Table 2. Number of active pigeon guillemot nestsat five colonies on the western side of Naked Island, Prince William Sound, Alaska, before and after T/V the Exxon Valdez oil spill. Dashes indicate no surveys were conducted.
Ye ar Nomad ThumbNomad Year RowTotal Tuft Hook Before spill 1978 I 11 9 12 19 58 19 79 8 1979 6 15 14 10 53 1980 5 4 10 8 6 33 1 981 4 1981 5 7 ______After spill 1989 3 4 I 2 5 21 1990 4 3 13 2 12 34 1994 5 2 3 29"11 10 "This value includes nests from both IHook and Hook I1 colonies. Note: Data from all years except 1994 from Table 2 (Oakley and Kuletz 1994).
30 Table 3. Growth rates of pigeon guillemot chicks raised at Naked Island, Prince William Sound, Alaska, before and after the T/V Exxon Valdez oil spill.
Mean Growth MinimumMaximum Number of RateStandard Growth Growth Ye ar ChicksYear (g/d)a Error Rate(g/d) Rate(g/d) Before spill 1 978 15 19.6 15 1978 7.4 1.4 31.7 1 979 16 23.8 16 1979 1.2 17.1 32.0 1980 bl 19.0 __ __ -_ 1 981 11 19.2 11 1981 1.8 11.4 34.3 After spill 1989 5 18.1 2.523.4 11.5 1990 12 16.6 1.2 10.1 23.6 1994 10 15.7 2.1 5.0 29.0 1 994 6 20.3 6 1994 1.4 15.0 23.5 (JI)‘ “Mean numberof grams gained per day during the linear growth phase, which is the period between8 and 18 days after hatching (Koelink 1972). bFew chicks were measured in 1980 becauseof loss of nests from netting and tagging of adults for foraging studies (Kuletz1983). ‘(JI) indicates data is from Jackpot Island. Note: Data from all years except 1994 from Table 14 (Oakley and Kuletz 1994).
31 Tab1.e 4. Fledging weights" of pigeon guillemot chicks raised at Naked Island, Prince WilliamSound, Alaska, before and after the T/V Exxon Valdez oil spill.
Mean Fledging Number of Weight StandardMinimum Maximum Ye ar ChicksYear (9)- Weight(g)Error Weight(g) Before spill 1978 29 467 9 467 29 1978 542 291 1979 17 506 12590 427 517 52 466 569 1980466 52 b2 517 1981 13 428 29 202 546 202 29 428 13 1981 After spill 1989 10 507 16570 420 1990 13 438 16 310 510 1994 17 453 13525 357 508 9 440 585 4401994 9 17 508 (JI)' "The last weight obtained from a chick that was measured within one week of fledging. 'Few chicks were measured in 1980 becauseof loss of nests from netting and tagging of adults for foraging studies (Kuletz 1983). '(JI) indicates data is from Jackpot Island. Note: Data from all years except 1994 from Table13 (Oakley and Kuletz 1994).
32 This page intentionally left blank. Figure 1. Locations (dots) investigated during the 1994 field season as potential study sites for pigeon guillemots breeding in Prince William Sound, Alaska.
34 r Figure 2. The Naked Island complex, comprising Naked, Peak, Storey, Smith, and Little Smith Islands,most of which were censused for pigeon guillemots by standardized methods in 1978-1980 and in 1989-1994.
36 Little Smith I "....v ethI. Figure 3. Locations of pigeon guillemot colonies (crosses) on Naked Island that were monitored during the 1994 field season.
38
Figure 4. Attendance patterns of pigeon guillemots at two colonies on Naked Island (13-14June 1994) during the incubationperiod. Times of high (H)and low (L) tides are indicated by letters at top of graph.
40 COLONYATTENDANCE NAKED I., 13-14 JUNE 1994
1 *TUFT + ROW
i ! j
I ! 1 i! -If! 16 17. 3 20 21 22 23 : I TIME OF DAY
41 Figure 5. Attendance patterns of pigeon guillemots at two colonies on Naked Island (5-6 July 1994) during the early hatchingperiod Times of high (H) and low (L) tides are indicatedby letters at top of graph.
42 COLONYATTENDANCE NAKED I., 5-6 JULY 1994 30 scn 25 45 u NOMAD ROW -4 - 3 (3 20 u. 0 15 W m E 10 1 3 z15 a 1 I 0 10 11 12 13 14 15 ' 19 20 21 22 23 24 01 02 03 04 05 06 07 08 09 TIME OF DAY
43 Figure 6. Nesting chronology of pigeon guillemots on Naked Island during the 1994 breeding season. Each bar represents one week beginning on the date below it. Only those nests followed from incubation through fledging were used to construct the chronology.
44 NAKED 1. NESTING CHRONOLOGY -- 1994
25 1median = 4 Junmedian =Jul median3 = 14Aug - 34 n n = 34 = 33 n=18
Days in Nest mean = 39.2 range = 30 - 48
5/155/22 5/29 6/05 611 2 6/196/26 7/03 7/10 711 7 7/247/31 8/07 8/14 8/21 DATE
45 Figure 7. Nesting chronology of pigeon guillemots on Jackpot Island during the 1994 breeding season. Each bar represents one week beginning on the date below it. Only those nests followed from incubation through fledging were used to construct the chronology.
46 JACKPOT 1. NESTING CHRONOLOGY -- 1994 30 median = 3 Jun median = 3 Jul median = 9 Aug n = 37 n = 37 n = 27 25
20 >. Days inNest 0 mean = 36.4 z range = 30 - 47 15 U W U U 10
5
0 1 - 511 5 5/225/29 6/05 6116/19 2 6/26 7/037/10 711 7 7/247/318/07 8/14 8/21 DATE
47 Figure 8. Comparison of In body mass vs. In wing length for chicks raised on Naked and Jackpot Islands. Second order polynomial equations for best-fit curves are shown.
48 PIGEON GUILLEMOTGROWTH Comparison between Islands -- 1994
0 Naked Island 6 Jackpot Island
~ ~
~ ~
.
0 OA Island: Naked OA . OA OA In(y) = - 3.932 + 3.554 In(x) - 0.309 (ln(x))"2 OA OA OA 144 observations on37 chicks from24 nests OA OA OA Jackpot Island: --O A A In(y) = - 6.548 + 4.660 In(x) - 0.422 (ln(x))"2 A A chicks95 31observations on nestsfrom 17
3.5 J 3 3.5 5 4 4.5 In wing length (mm)
49 Figure 9. Temporal distribution offood deliveries by adult pigeon guillemots at several colonies on Naked Island during the 1994 breeding season.
50 TEMPORALDISTRIBUTION OF DELIVERIES NAKED ISLAND -- 1994
< 06 07 08 09 10 11 12 13 14 15 16 17 18 19 20 21 > TIME OF DAY
51 Figure 10. Temporal distribution of food deliveries by adult pigeon guillemots at Jackpot Island during the 1994 breeding season.
52 TEMPORALDISTRIBUTION OF DELIVERIES JACKPOT ISLAND -- 1994 20 , tn W E 15 -- >W i W n -- u 10 0 a W rn z 5-- z3
n, < 06 07 08 09 10 11 12 13 14 15 16 17 18 19 20 21 > TIME OF DAY
53 Figure 11. Temporal distribution of food deliveries (inverted triangles) to each of four guillemot nests at Tuft colony on Naked Island (3 August 1994). Notations above lines indicate total number of fish (0 delivered, delivery rate (fi),and number of chicks (c) in the nest on that day. A "?" indicates the number of chicks in the nest on that day was not known and, if followed by a "(2)", that two chicks hatched in that nest. Two deliveries occurring within a three-minute period are indicated by a "+" above a triangle.
54 FOODDELIVERIES Tuft, Naked 1. -- 3 Aug 1994
T1
T8
T9
21f1.171h ?c (2) T10 - - - --
0500 2300 TIME OF DAY
55 Figure 12. Temporal distribution offood deliveries (inverted triangles) to each of three guillemot nests at Nomad colony on Naked Island (7 August 1994). Notations above lines indicate total number of fish (0 delivered, delivery rate (fi),and number of chicks (c) in the nest on that day. A "?" indicates the number of chicks in the nest on that day was not known and, if followed by a "(2)", that two chicks hatched in that nest. Two deliveries occurring within a three-minute period are indicated by a "+" above a triangle.
56 FOOD DELIVERIES Nomad, Naked 1. -- 7 Aug 1994
N1
1.171h 1 c ++ N3 -7 r;r rr -.-
N5
0500 2300 TIME OF DAY
57 Figure 13. Temporal distribution of food deliveries (inverted triangles) to each of four guillemot nests on Jackpot Island (6 August 1994). Notations above lines indicate total number of fish (0 delivered, delivery rate (fi),and number of chicks (c) in the nest on that day. A "?" indicates the number of chicks in the neston that day was not known and, if followed by a "(2)", that two chicks hatched in that nest. Two deliveries occurring within a three- minute period are indicated by a "+" above a triangle.
58 FOOD DELIVERIES Jackpot 1. -- 6 Aug 1994
J10
J11
J13
f 0.891 h ? c (2) J29 v
0500 2300 TIME OF DAY
59 Figure 14. Bathymetry of the Naked Island area. Note the broad expanse of shallow water extending aroundmost parts of Naked, Peak, and Storey Islands.
60 I Bathymetric Depth Classification
0....., :. < 20 Meters 60 - 80 Meters @ 120. 200 Meters ~ @ 20 - 40 Meters @ 80 - 100 Meters 0 > 200 Meters Figure 15. Bathymetry of the Jackpot Island area. Note the deep water surrounding Jackpot Island and shallower water near the mouths of Jackpot Bay and Icy Bay.
62
Figure 16. Comparison of prey types and relative proportions of each delivered by adult pigeon guillemots at several colonies on Naked Island (16 observations) and at Jackpot Island (3 observations). All observations were made between 17 July and 12 August 1994.
64 PREYTYPES 81 PROPORTIONSDELIVERED BY PIGEON GUILLEMOTS herring I smelt (1.2%)~ unknown (25.4%) NAKED 1. n = 1061
sand lance (7.9%
All observations made between 17 July and 12 August 1994.NI = 16 obs, JI = 3 obs
-unknown (24.2%)
flatfish (1.8%) scubins (2.6%)
UlUS ( 10.370) (6.3%)iprickledacks I cnnels(1 1.7%) Lsand lance (0.8%) APPENDIX
Oakley and Kuletz (1994) included several tables in their report, eachof which relates to some measure of pigeon guillemot productivity. These tables have been reproduced here with only minor modifications, and updated with the appropriate values for 1994. Although the original tables contained information from Naked I. only, data from Jackpot I. have been included for comparative purposes.
Table Al. Mean clutch size (mean number of eggs per nest found during the egg stage) of pigeon guillemots at Naked Island, Prince William Sound, Alaska, before and after the TIV Exxon Valdez oil spill ...... 67
Table A2. Mean number of chicks hatched per nest found in the egg stage of pigeon guillemots at Naked Island, Prince William Sound, Alaska, before and after theTIV Exxon Valdez oil spill ...... 68
Table A3. Mean number of chicks fledged per nest found in theegg or chick stage of pigeon guillemots at Naked Island, Prince William Sound, Alaska, before and after the TIV Exxon Valdez oil spill ...... 69
Table A4. Mean number of chicks fledged per nest found in the egg stage of pigeon guillemots at Naked Island, Prince William Sound, Alaska, before and after theTIV Exxon Valdez oil spill ...... 70
Table A5. Causes of nesting failureof pigeon guillemots at Naked Island, Prince William Sound, Alaska, before and after the TIV Exxon Vuldez oil spill ...... 71
66 Table Al. Mean clutch size (mean numberof eggs per nest found during the eggstage)a of pigeon guillemots at Naked Island, Prince William Sound, Alaska, before and after the T/V Exxon Valdez oil spill.
Number of Meanclutch Standard nests size error Year size nests Before spill 1978 13 1.54 0.14
1979 33 1.85 0.06
1980 27 1.78 27 1980 0.08
1981 22 1.59 22 1981 0.11
1984 7 1.86 0.14
After spill 1989 7 1.57 0.20
1990 27 1990 1.78 0.10
1994 b2 1994 3 1.70 0.10
1994(JI)' 24 1.92 0.09 "The mean number of eggs per nest found during the egg stage is what Oakley and Kuletz(1994) refer to as mean clutch size; it is equivalent to mean clutch size as used in this report. bThis value includes23 clutches from 22 nests. '(JI) indicates data from Jackpot Island. Note: Data from all years except 1994 from Table8 (Oakley and Kuletz 1994).
67 Table A2. Mean number of chicks hatched per nest found in the egg stage“ of pigeon guillemots at NakedIsland, Prince William Sound, Alaska, before and after T\V the Exxon Valdez oil spill.
Year Number of Mean number of Standard nests chicks hatched error Before spill 1978 9 1.22 0.28
1979 32 1979 1.34 0.15
1980 bl. 20 05 0.20
1981 21 1981 1.14 0.17
1984 7 1984 0.37 1.43
After spill 1989 7 1.43 0.30
1990 25 1.28 25 1990 0.16
1994 ‘23 1.52 0.15
1994 24 1.54 24 1994 0.17 “The mean number of chicks hatched per nest found in the egg stage is what Oakley and Kuietz (1994) refer to as hatching success rate; it is NOT equivalent to hatching successas used in this report. bNests of birds subjected to netting and tagging for foraging studies were not included, but reproduction of neighboring pairs at colonies where netting and tagging occurred might have been affected (Kuletz 1983). ‘This value includes 23 clutches from 22 nests. d(JI) indicates data is from Jackpot Island. Note: Data from all years except 1994 from Table 9 (Oakley and Kuletz 1994).
68 Table A3. Mean number of chicks fledged per nest found in the egg or chickstagea of pigeon guillemots at Naked Island, Prince WilliamSound, Alaska, before andafter the T/V Exxon Valdez oil spill.
NumberofMean number of Standard Year nests chicks fledged error fledged chicks nests Year Before spill 1978 30 1978 1.60 0.10 1979 23 1.40 0.15 1980 12 bl. 00 0.25 1981 17 1.06 0.18 1984 6 1984 1.17 0.31 After spill 1989 14 1989 0.93 0.22 1990 29 1990 0.79 0.14 1994 ‘35 0.86 0.13 1994 (JI) 28 1.25 0.15 “The mean numberof chicks fledged per nest found in egg the or chick stage is what Oakley and Kuletz (1994) refer to as fledging success rate; it is NOT equivalent to fledging success as used in this report. bNests of birds subjected to netting and tagging for foraging studies were not included, but reproductionof neighboring pairs at colonies where netting and tagging occurred might have been affected (Kuletz 1983). ‘This value includes35 clutches from 34nests. d(JI) indicates data is from Jackpot Island. Note: Data from all years except 1994 from Table10 (Oakley and Kuletz 1994).
69 Table A4. Mean number of chicks fledged per nest found in the egg stage“ of pigeon guillemots at NakedIsland, Prince William Sound, Alaska, before and after T/V the Exxon Valdez oil spill.
Number of Mean number of Standard Year nests chicks fledged chicks nests Year error Before spill 1978 8 1.25 0.20 1.25 8 1978 1979 30 1.13 0.13 1.13 30 1979 1980 18 1980 61 0.20 1981 19 1981 0.74 0.17 1984 7 1.00 0.28 After spill 1989 0.34 6 0.50 1990 24 0.68 0.16 1994 0.17 ‘23 0.78 1994 (JI)d 24 1.17 0.17 “The mean number of chicks fledged per nest found in the egg stage is what Oakley and Kuletz (1994) refer to as nesting success rate; it is NOT equivalentto nesting success as used in this report. bNests of birds subjected to netting and tagging for foraging studies were not included, but reproduction of neighboring pairs at colonies where netting and tagging occurred might have been affected (Kuletz 1983). ‘This value includes 23 clutches from nests. 22 d(JI) indicates data is from Jackpot Island. Note: Data for all years except 1994 from Table 11 (Oakley and Kuletz 1994).
70 Table A5. Causesof nesting failure of pigeon guillemots at Naked Island, Prince William Sound, Alaska, before and afterthe T/V Exxon Valdez oil spill. Number of Nests in whichat least one egg failed to hatch or at least onechick failed to fledge. Young Starva- Number of Unhatched Chick Predation Predation tion or Unknown Year NestsEgg" Deathb Egg' of of Chick' ExposureReason
Betore spill 1978 32 4 0 0 0 0 2 1979 30 6 1 2 1 1 4 1980 19 1980 5 0 2 0 4 2 1981 22 6 1 2 0 6 0 After spill 1989 15 3 1 1 4 0 0 1990 38 2 1 3 4 1 5 1994 23d 1 4 1 1 2 7 1994(JI)' 24 5 2 0 0 0 6 "Includes eggs which failedto hatch due to infertility, embryodeath, or nest desertion. bRefers to chicks, less than one weekold, dying in the nest for no apparentreason. 'In previous years, predation was rarely witnessed; most instances of nesting failure attributed to predation were presumed based on disappearanceof eggs or chicks. In 1994, predation was never witnessed,but conclusive evidence of predation was found. dThis value includes 23 clutches from22 nests. '(JI) indicates data is from Jackpot Island. Note: Data from all years except 1994 from Table12 (Oakley and Kuletz 1994).